Research progress of the heritable melanism mutation of insects
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摘要:
体色变异在昆虫适应复杂环境中发挥着十分重要的作用，黑化是一种最常见的体色多型现象。其中，许多黑化是可遗传的突变。不同突变体的黑化表型呈现阶段与部位及形成机制具有较大差异，其生物学特性变化也因种而异。许多昆虫都会自发产生黑化突变体，它们是探究黑化形成机制及基因多效性的理想研究材料。为了全面了解昆虫黑化突变体的主要类型及其产生机制。本文围绕呈现黑化表型的发育阶段及部位，黑化突变的遗传学基础、生理与分子机制及生物学特性变异等方面进行了综述，并对未来研究方向进行了探讨。
Abstract:Variation of body color plays important roles in the environmental adaption of insects, and melanism is the most common type of polymorphism. Among them, many melanism phenotypes are inheritable and called mutants. Different mutants display a wide range of melanism phenotypes. Moreover, the underlying molecular mechanisms and the consequent phenotypic alterations vary significantly among different mutants. Many insects possess spontaneously occurring melanism mutants, which serve as crucial materials for uncovering the mechanisms of melanism and the pleiotropy of melanism genes. In order to fully understand the melanism mutant type and the caused mechanisms. Here, we made a comprehensive review of the information related to melanism mutants, including development stage and region displayed melanism phenotype, physiological and molecular mechanisms, and variation of biological features. Additionally, we made a discussion for the future hot topics in this research field.
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Keywords:
- Melanism /
- spontaneous mutations /
- insect /
- molecular mechanism /
- pleiotropy
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体色对于生物的生存与进化有着十分重要的作用，是一种变异频率较高的性状（杨建华和徐安英，2017）。昆虫是地球上种类最多的动物类群（Vogel et al.，2019），其丰富多样的体色和斑纹在躲避敌害和适应环境变化方面具有十分重要的作用。黑化（Melanism）是昆虫最为普遍的体色变异现象（True，2003），最先由Edleston（1864）在桦尺蠖中观察到。True（2003）对黑化进行了全面定义：表皮呈现大部分或完全变黑的变异个体或群体现象。研究表明多种因子可引起昆虫的黑化，如温度、工业污染等，且部分黑化表型是可以稳定遗传的，即为突变体（江幸福和罗礼智，2007）。体色突变体的产生对解析色素沉积演化，体色变异的遗传基础和分子机制具有十分重要的作用（Protas and Patel，2008）。此外，仅在某些特定阶段呈现黑化的突变体还为解析昆虫的非偶联发育机制提供了良好材料（Saenko et al.，2012），相关突变位点对探究宏观与微观进化具有重要作用（Haag and True，2001）。同时，这些可见的表型差异还可用于种内及种间竞争研究（Park et al.，1945）。室内饲养环境虽相对稳定，但一些昆虫仍能自发产生黑化突变体，如家蚕就拥有20余种黑化突变品种（张彦等，2017）。
昆虫黑化一直是人们关注的热点，多位学者在前期已从不同角度对黑化现象及其机理进行了总结。朱福兴等（2007）重点关注了影响昆虫黑化的因素、色素形成机理、黑化突变的机理与遗传等。江幸福与罗礼智（2007）则从黑化定义、昆虫黑化现象、黑化的产生原因及类别、黑化的遗传及分子基础、黑化昆虫的生物学变异等方面进行了综述，尤其是对昆虫黑化的产生原因及类别、遗传学基础等进行了全面总结。高云等（2020）对黑色斑点与斑纹的形成和调节机制（以鳞翅目家蚕与虎凤蝶为代表）进行了总结。目前，针对黑化突变体的相关研究进展仅在家蚕中有较为系统的总结。张彦等（2017）分别从黑色素合成代谢途径与相关功能基因、几种主要黑化突变品系的研究、黑化突变对生物特性的影响等方面进行了综述；杨建华和徐安英（2017）重点介绍了家蚕突变体的鉴定与筛选及分子水平的相关研究进展等。本文将以昆虫主要自发黑化突变体为对象，对黑化表型呈现的发育阶段及部位、遗传学基础、生理与分子机制及其生物学特性变异等进行总结分析（详见网络版增强出版材料附表 1），旨在为进一步阐明昆虫丰富体色的形成机制奠定基础。
1. 呈现黑化表型的发育阶段及部位
在不同突变体中，表现黑化的发育阶段存在较大差异。其中，部分种类仅在某个发育时期呈现出黑化表型。成虫期黑化：鳞翅目黑化粘虫Mythimna separata（刘红兵和罗礼智，2004）及家蚕Bombyx mori中的黑蛾Bm（Futahashi et al.，2018）与暗化型mln-2（张凤林等，2009），鞘翅目杂拟谷盗Tribolium confusum中的ebony与McGill black（Park et al.，1945；Stanley and Slatis，1955）及具斑食蚜瓢虫Coleomegilla maculata，双翅目黑腹果蝇Drosophila melanogaster中的4个突变体（黑檀体ebony、黑体black、sable及黑条体ebonybsr）（Jacobsm，1960；Hodgetts，1972；钱远槐等，1994，1995；Dean et al.，2022）。特别地，家蚕从性黑蛾sml仅在雄性成虫中表现黑化（代方银等，2013），而四纹豆象Callosobruchus maculatus以雌性成虫表型最为明显（Breitenbecher，1921）。蛹期黑化：黑化甜菜夜蛾Spodoptera exigua SEM（刘子晶，2006）与家蚕中的黑蛹bp（Dai et al.，2015）。幼虫期黑化：家蚕中的黑缟pS与bd（Kawaguchi et al.，2007；刘丽丽等，2014）及黑化眼斑蝶Bicyclus anynana dl（Bear et al.，2010）。特别地，烟草天蛾Manduca sexta bl仅在末龄幼虫表现黑化（Safranek and Riddiford，1975）。此外，一些突变体在某两个发育阶段呈现黑化，主要为老熟蛹与成虫的黑化：棉铃虫Helicoverpa armigera的两个黑化突变体（马伟华，2005；何丽华等，2010）、杂拟谷盗中的negret（Neelam and Bhatia，1983）及绿豆象Callosobruchus chinensis中的sC-b（Kashiwagi and Utida，1972）。最后，还有少数突变体在生长发育的幼虫、蛹与成虫3个阶段均呈现黑化。例如，家蚕中的暗化型mln、浅暗化型及so突变（Futahashi，2008；Dai et al.，2010；张彦等，2018），黑化赤拟谷盗Tribolium castaneum（Sokoloff et al.，1960）与异色瓢虫Harmonia axyridis ml突变体（李明凌等，2022；Sun et al.，2025）。特别地，家蚕so突变的黑化在蛹期表现最为明显（Futahashi，2008）。
2. 黑化自发突变的遗传学基础
昆虫自发黑化突变体的遗传方式多样，包括孟德尔式遗传与非孟德尔式遗传两类。其中，遵循孟德尔式遗传的大部分种类为常染色体上的隐性突变，约占已报道种类的53.3%（16/30）。常染色体隐性遗传：鳞翅目昆虫中的绝大数黑化突变体及鞘翅目中的杂拟谷盗ebony（Park et al.，1945）、异色瓢虫ml、绿豆象sC-b、鹰嘴豆象与烟草甲（详见网络版增强出版材料附表 1）；家蚕从性黑蛾sml还与性别关联，仅在雄性成虫中表现黑化。常染色体显性遗传：家蚕中的黑缟（pS）（刘丽丽等，2014）、杂拟谷盗中的McGill black（Stanley and Slatis，1955）及黑化四纹豆象（Breitenbecher，1921）。常染色体上的不完全显性（半显性）遗传：甜菜夜蛾黑蛹SEM（刘子晶，2006；刘向阳，2007）、黑化赤拟谷盗（Sokoloff et al.，1960）及具斑食蚜瓢虫（Allen，2016）。此外，黑化烟草天蛾bl为特殊的雌性连锁的单基因突变，并受其他基因的修饰（Truman et al.，1973；Safranek and Riddiford，1975）；杂拟谷盗中的negret为雄性连锁的隐性突变（Neelam and Bhatia，1983）。目前，已报道的非孟德尔式遗传的典型代表为黑腹果蝇中的黑条体ebonybsr，黑条体性状在杂交后代中表现很不稳定（钱远槐等，1994，1995）。
3. 黑化自发突变的生理与分子机制
通过多年研究，昆虫黑色素的合成通路已较为清楚，其主要步骤如下所示。首先，在表皮细胞中，由TH基因编码的酪氨酸羟化酶（Tyrosine hydroxylase，TH）将酪氨酸转化为多巴（Dopa），多巴由多巴脱羧酶（Dopa decarboxylase，DDC）转化为多巴胺（Dopamine）。多巴胺和多巴是最重要的两类色素前体物质，它们在Yellow家族基因和Lac2的作用下可分别转化为多巴胺黑色素或者多巴黑色素。随后，合成的色素前体从表皮细胞中分泌出来掺入表皮，形成黑色斑纹和斑点模式。除此之外，多巴胺还有两条代谢途径。一是，多巴胺可与β-丙氨酸（β-alanine）结合由ebony转化为N-β-丙氨酰多巴胺（N-β-alanyldopamine，NBAD），参与表皮鞣化，且使表皮呈现黄色或者淡黄色，这一过程可以被tan基因编码的N-β-丙酰多巴胺水解酶（NBAD hydrolase）逆转。在此过程中的β-丙氨酸（β-alanine）是由天冬氨酸在天冬氨酸脱羧酶（Aspartate decarboxylase，ADC）的作用下转化而来的。二是，多巴胺还可由芳基烷基胺-N-乙酰基转移酶（Arylalkylamine N-acetyltransferase，AA-NAT）转化为N-乙酰基多巴胺（N-acetyl dopamine，NADA），作为无色硬化蛋白的前体参与表皮硬化（Noh et al.，2016；高云等，2020）（图 1）。此外，研究发现表皮蛋白基因与黑化具有密切关联，家蚕幼虫表皮蛋白基因的表达量与黑化程度呈正相关（Qiao et al.，2020）。不同黑化突变体的形成机制具有较大差异，围绕黑色素代谢途径中多巴胺的代谢通路可将主要黑化突变体的生理与分子机制进行如下归类。
注：绿色椭圆框代表色素合成通路的关键基因，黑色、黄色和白色框代表色素前体沉积到表皮的颜色。Fig. 1 Pathway for Melanin synthesis in insects (Modified from Noh et al., 2016 and Gao et al., 2020)Note: The green oval boxes represented the key genes of the pigment synthesis pathway, and the black, yellow and white boxes represented the color of the pigment precursor deposited on the epidermis, respectively.
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首先，由于多巴胺合成异常引起其过量积累而产生黑化突变。其中，甜菜夜蛾蛹期的黑化突变是由于TH与DDC过表达引起的（Liu et al.，2015a）。黑化烟草天蛾体内DDC也是过量表达的（Hiruma and Riddiford，2009），且其过量表达是受幼虫体内保幼激素（JH）含量下降所引起（Truman et al.，1973；Safranek and Riddiford，1975；Hiruma and Riddiford，2009）。家蚕黑缟（pS）与黑化赤拟谷盗产生的生理基础也是多巴胺含量增加，但其具体调控机制尚不明确（Lomakin et al.，2010；张彦等，2021）。
其次，由于多巴胺代谢异常引起其过量积累而产生黑化突变。目前，许多黑化突变体都表现为β-丙氨酸不足及与多巴胺反应的异常，从而积累大量多巴胺。β-丙氨酸合成异常：天冬氨酸脱羧酶基因ADC（合成β-丙氨酸）突变是产生家蚕黑蛹突变（bp）与黑腹果蝇黑体突变（black）的重要原因。在家蚕bp突变体中，一个493 bp的SINE-like转座子插入ADC转录起始位点上游~2.2 kb引起其转录水平的显著降低。上述机制通过向bp突变体中注入β-丙氨酸能将突变表型恢复为野生型得以进一步证实（Dai et al.，2015）。β-丙氨酸代谢异常：在黑腹果蝇黑檀体突变（ebony）中，β-丙氨酸转氨酶活性增强使得β-丙氨酸的代谢增强（Wittkopp et al.，2002；艾炎军和曾庆韬，2010）。β-丙氨酸与多巴胺反应异常：黑化赤拟谷盗的产生是由于β-丙氨酸缺乏N端酰化无法与多巴胺发生合成反应（Roseland et al.，1987；Lomakin et al.，2010）。此外，家蚕的so突变是由于Ebony基因开放阅读框的大片段缺失（Futahashi，2008）；黑腹果蝇黑条体ebonybsr是由于Ebony基因5'端UTR区和第1个外显子部分缺失（金珊和曾庆韬，2010）；在黑腹果蝇sable突变中，mdg4插入Yippee基因的5'端UTR区，也引起多巴胺合成NBAD的异常（Dean et al.，2022）；在家蚕暗化型突变（mln）中，由于AANAT的功能失调使得多巴胺无法转换为NADA，是昆虫中第一种涉及AANAT的黑化突变体（Dai et al.，2010；Zhan et al.，2010；Qiao et al.，2012）。
最后，由于黑色素合成步骤异常引起产物过量积累而产生黑化突变。棉铃虫蛹黑型突变体与酚氧化酶活力突增有关，其黑化突变体中的酚氧化酶活力是野生型的234倍（马伟华，2005）。家蚕从性黑蛾突变（sml）是由yellow与laccase2基因的上调表达引起（代方银等，2013；He et al.，2016）。此外，家蚕黑化突变bd是由于表皮蛋白基因上调表达引起（Kawaguchi et al.，2007；Wu et al.，2016）。
4. 黑化自发突变的生物学特性变异
昆虫黑色素合成的起始物质—酪氨酸只能从食物中获取，并且在合成中需要大量氮气或蛋白质来提供氮原子（True，2003；Gonzalez-Santoyo and Cordoba-Aguilar，2012）。因此，一般认为黑化会影响昆虫的适合度，即“黑化导致适合度降低”假说。目前，在黑化突变体中已测定的生物学特性主要涉及生长发育、寿命、个体大小、生殖与视觉等。
首先，黑化突变引起部分个体适合度的降低。在生长发育方面，黑化棉铃虫JBM发育速度变慢，成虫个体变小（Ma et al.，2008）；家蚕暗化蛾茧重略有降低（杨明观等，2001）；烟草天蛾bl发育速率、存活率及体重均降低（Safranek et al.，1975）；杂拟谷盗McGill black与negret发育时间延长，且negret在蛹期与成虫羽化期的存活率下降（Edwards，1958；Neelam and Bhatia，1983）。特别地，黑缟（pS）具有纯合致死特性（刘丽丽等，2014；张彦等，2017）。在生殖方面，黑化棉铃虫JBM、烟草天蛾bl、眼斑蝶dl、杂拟谷盗ebony与negret及绿豆象sC-b均表现为生殖力降低（Park et al.，1945；Kashiwagi and Utida，1972；Safranek and Riddiford，1975；Neelam and Bhatia，1983；Ma et al.，2008；Bear et al.，2010）。此外，黑化棉铃虫JBM还表现出交配率降低（Ma et al.，2008）；黑腹果蝇黑条体ebonybsr表现为求偶动作减少（艾炎军和曾庆韬，2010）；家蚕黑化突变bd表现为雌性不育（Kawaguchi et al.，2007）。在视觉行为方面，黑腹果蝇黑檀体ebony与黑体black对视觉信号反应异常（Jacobsm，1960；Hodgetts，1972；Phillips et al.，2005）。在表皮特性方面，黑化赤拟谷盗的鞘翅较野生型具有更高的粘滞能量耗散程度（Lomakin et al.，2010）。
其次，黑化突变引起个体适合度的改善。黑化粘虫幼虫期和蛹期的发育时间均缩短，存活率提高；成虫产卵前期缩短且产卵量增加（Jiang et al.，2007）。甜菜夜蛾SEM突变的幼虫发育速度加快，蛹重增加；成虫交配时间延长且生殖力提高（Liu et al.，2015b）。家蚕从性黑蛾sml寿命延长并具更大活力（代方银等，2013）；异色瓢虫黑化突变ml取食人工饲料后的成虫畸形率降低（李明凌等，2022）；黑化四纹豆象的雌虫个体增大（Breitenbecher，1921）；果蝇黑化突变体ebony的体温升高（Freoa et al.，2023）。
除此之外，少数种类的黑化突变对个体适合度无明显不利影响。鹰嘴豆象黑化突变体具有与野生型相似的繁殖力、寿命与卵孵化率（Numajiri et al.，2017）；烟草甲黑化突变体的交配行为与生活史均未发生改变（Omae et al.，2012）。
5. 结语和展望
目前已报道的昆虫自发黑化突变体主要集中于鳞翅目、鞘翅目与双翅目中，且以鳞翅目居多，这些种类大多为重要的农林害虫及经济昆虫，但在不同种类研究中的关注点有所差异。其中，家蚕与果蝇突变体主要用于阐释黑化分子机制。随着对一系列突变体的研究，二者的黑色素代谢通路越来越清晰，它们也因此成为研究色素沉积的模式昆虫（Dai et al.，2015；Dean et al.，2022）。其他一些农林害虫则重点关注其生物学特性变异，并在甜菜夜蛾与烟草天蛾等少数种类中进行了分子机制方面的探究。昆虫体色和斑纹多样性是研究发育生物学、自然选择和形态性状进化的良好对象（高云等，2020）。围绕这些黑化突变体，今后应从以下方面开展研究。
首先，不同突变体的黑化部位及发生时期都存在较大差异，因此每一种突变体都具有特异的黑化机制（Wang et al.，2022）。研究表明，桦尺蠖自然种群的黑化突变与目前所鉴定的16个黑色素合成关键基因均无关，其突变机制是一个串联重复的大转座子插入到了基因cortex的第一个内含子区（van't Hof et al.，2016）；家蚕mln是第一种涉及AANAT基因的昆虫黑化突变体（Dai et al.，2010）。室内饲养种群的黑化突变体大都是自发产生的，针对这些突变体分子机制的系统解析有助于阐明昆虫黑化途径的易突变位点。同时，基于种间的异同分析可揭示这些易突变位点的进化意义。
其次，基因多效性是生物体中普遍存在的一种现象，即某一基因发生突变往往会导致其他性状的改变（Massey et al.，2019）。近年来，昆虫体色基因的多效性已成为研究热点（李明凌等，2022）。目前，有关自发黑化突变体生物学特性变异研究所涉及的参数在不同种类中存在较大差异，如求偶与视觉行为主要集中于黑腹果蝇中。上述研究参数的不统一就导致突变基因的多效性无法进行系统分析。在未来，研究人员应围绕突变体建立标准一致的生物学特性变异评价体系。
最后，昆虫作为变温动物，其响应环境变化的体色变异特性近年来受到了广泛关注（Schilthuizen and Kellermann，2014；MacLean et al.，2018）。通常黑化程度高的个体由于体温上升较快而在低温环境下生存有利（Clusella-Trullas and Nielsen，2020），许多昆虫的表皮黑化程度也因此具有温度可塑性（Zhang et al.，2021），但目前围绕自发黑化突变体响应温度变化的体色变化规律及其温度适应能力研究还较少，仅局限于果蝇（Freoa et al.，2023）及异色瓢虫（Sun et al.，2025）等少数物种中。针对这些黑化突变体的环境适应机制研究，可为进一步明确昆虫体色的变化规律及开展有益种类的饲养与应用奠定良好基础。
附录：附表 1 昆虫黑化突变体及其形成机制与生物学特性变异
详细数据见网络版增强出版材料（http://hjkcxb.alljournals.net）
附表 1 昆虫黑化突变体及其形成机制与生物学特性变异Appendix Table 1 Insect melanism mutants and the mechanisms and changing of biological characteristics目
Order科
Family种名
Species名称及黑化阶段
Name and melanism stages遗传学规律
Genetic basis生物学特性变异
Changing of biological characteristics生理与分子机制
Physiological and molecular mechanism参考文献
References鳞翅目
Lepidoptera夜蛾科
Noctuidae棉铃虫
Helicoverpa armigera HübnerJBM
老熟蛹和成虫
Mature pupa and adultAS- 发育速度变慢, 成虫个体变小, 交配率降低, 生殖力下降
Decreased development rate, reduced adult body size, decreased mating rate and reproductive output酚氧化酶活力突增
Significantly increased activity of phenoloxidase能量代谢与细胞骨架组分蛋白的含量改变Changes of energy metabolism and cytoskeletal protein contents马伟华, 2005;Wang et al., 2005;Ma et al., 2008
何丽华等, 2010;何丽华等, 2012—
老熟蛹和成虫
Mature pupa and adultAS- 粘虫
Mythimna separate (Walker)—
成虫
AdultAS- 幼虫和蛹发育加快, 存活率提高; 产卵前期缩短且产卵量增加
Accelerated development of larva and pupa, increased survival, reduced pre-oviposition period and increased oviposition刘红兵和罗礼智, 2004;Jiang et al., 2007 甜菜夜蛾
Spodoptera exigua (Hübner)SEM
蛹
PupaAS± 发育速度加快, 蛹重增加, 交配时间延长, 生殖力提高
Accelerated development, increased pupal weight and mating period, increased reproductive capabilityTH与DDC过表达
Overexpression of TH and DDC刘子晶, 2006;刘向阳, 2007;刘思思, 2015;Liu et al., 2015a, 2015b 蚕蛾科
Bombicidae家蚕
Bombyx mori L.暗化型mln
幼虫、老熟蛹及成虫
Larva, mature pupa and adultAS- AANAT的功能失调及与儿茶酚胺代谢的交互作用
Dysfunction of AANAT and its interactive with catecholamineDai et al., 2010;Zhan et al., 2010;Qiao et al., 2012 浅暗化型Lightmelanism
幼虫、老熟蛹及成虫Larva, mature pupa and adultAS- 多巴、多巴胺含量改变 Changes of dopa and dopamine content 张彦等，2018，2021 黑蛾 Bm
成虫 Adult
soAS-Banno et al.，2005 幼虫、蛹及成虫, 以蛹最明显Larva, pupa and adult, but predominant in pupa ebony 基因开放阅读框的大片段缺 失 Large fragment deletion of the ORF of ebony Futahashi，2008 黑蛹bp
蛹PupaADC 基因起始位点调控序列插入转 座子 Insertion of transposons in the initial regulatory sequence of gene ADC Dai et al.，2015 从性黑蛾sml
雄性成虫Male adultAS-, 与性别关联Sex-related 寿命延长并具有 更高活力 Prolonged the lifespan and expressed higher activity yellow 与 laccase2 基因的上调表达 Up-regulated expression of gene yellow and laccase2 代方银等，2013；
He et al.，2016暗化蛾Melanism moth
幼虫及成虫Larva and adult mln-2AS- 茧重略有减轻 A slight decrease in weight of cocoon 杨明观等，2001 成虫Adult AS- 张凤林等，2009 黑缟(ps)
幼虫LarvaAS- 酪氨酸向多巴的转化速度提高 Accelerated transformation of tyrosine to dopa 刘丽丽等，2014；
张彦等，2017bd
幼虫LarvaAS- 雌性不育Infertility of female 表皮蛋白基因上调表达Up-regulated expression of cuticular protein gene Kawaguchi et al., 2007;Wu et al., 2016 天蛾科Sphingidae 烟草天蛾Manduca sexta L. bl
末龄幼虫Last-instar larva雌性连锁单基因, 并受其他基因修饰Female-linked single gene and modified by other genes 发育速率、体重、存活率及生殖力降低Decreased development rate, body weight, survival and fecundity 保幼激素含量不足, DDC基因上调表达Insufficient juvenile hormone content, and up-regulated expression of DDC Truman et al., 1973;Safranek and Riddiford, 1975; Hiruma and Riddiford, 2009 蛱蝶科Nymphalidae 眼斑蝶Bicyclus anynana (Butler) dl
幼虫LarvaAS- 生殖力降低Decreased fecundity Bear et al., 2010 赤拟谷盗Tribolium castaneum (Herbst) —
幼虫、蛹及成虫Larva, pupa and adultAS± 鞘翅的机械特性发生改变Changes of the mechanical characteristics of elytra 多巴胺含量增加Increased dopamine content Sokoloff et al., 1960;Kramer et al., 1984;Roseland et al., 1987;Lomakin et al., 2010 ebony
成虫AdultAS- 产卵量下降Decreased oviposition Park et al., 1945 鞘翅目Coleoptera 拟步甲科Tenebrionidae McGill black成虫Adult AS+ 发育时间延长Prolonged development time Stanley and Slatis, 1955;Edwards, 1958 杂拟谷盗Tribolium confusum Duval negret
老熟蛹及雄成虫Mature pupa and male adult雄性连锁的隐性调控Male-linked recessive regulation 发育速率变缓, 生殖力下降, 蛹期与成虫羽化期的存活率下降Slowdown of the development rate, decreased fecundity, decreased survival rates of the pupae and newly emerged adults Neelam and Bhatia, 1983 瓢甲科Coccinellidae 具斑食蚜瓢虫Coleomegilla maculata (DeGeer) 成虫鞘翅Elytra of adult AS± Allen, 2016 异色瓢虫 ml 羽化成虫畸形率降低; 鞘翅类胡萝卜素浓度升高; 抗紫外能力增强Decreased the malformation rate of newly emerged adults; Increased elytra carotenoid concentration; increased UV-resistance 李明凌等, 2022;李媛等, 2025;Sun et al., 2025 Harmonia axyridis (Pallas) 幼虫、蛹和成虫Larva, pupa and adult AS- 绿豆象Callosobruchus chinensis (L.) sC-b 繁殖力与生育力降低Decreased fecundity and fertility Kashiwagi and Utida, 1972 老熟蛹及成虫Mature pupa and adult AS- 豆象科
Bruchidae四纹豆象Callosobruchus maculatus (Fabricius) 成虫, 雌虫最明显Adult and predominant in female AS+ 雌虫个体大一些Increased body size of female adults Breitenbecher, 1921;Eady, 1991;Mano and Toquenaga, 2011 鹰嘴豆象Callosobruchus analis (Fabricius) 成虫
AdultAS- 适合度无影响
No significant effects on fitnessNumajiri et al., 2017 窃蠢科
Anobiidae烟草甲Lasioderma serricorne (Fabricius) 成虫
AdultAS- 交配行为与生活史均未发生改变No changes of mating behavior and life history Omae et al., 2012 黑檀体 ebony
成虫Adult交尾潜伏期增加, 交配及视觉行为异常Increased mating latent period, and exhibited abnormal mating and vision behavior β-丙氨酸代谢发生缺陷Defect in β-alanine metabolism Jacobsm, 1960;Hodgetts, 1972;Wittkopp et al., 2002;Kohn and Wittkopp, 2007 双翅目
Diptera果蝇科
Drosophilidae黑腹果蝇
Drosophila melanogaster Meigen黑体black
成虫Adult对视觉信号的反应异常Abnormal response to visible signals ADC基因突变Mutation of ADC gene
mdg4插入Yippee基因的5'端UTR区Insertion of mdg4 in 5' UTR region of Yippee geneHodgetts, 1972;Phillips et al., 2005 sable
成虫Adult√ Dean et al., 2022 黑条体ebonybsr
成虫Adult非孟德尔式遗传
Non-Mendelian inheritance求偶动作减少、体温升高Reduced courtship behavior, and increased body temperature ebony基因5'端UTR区和第1个外显子部分缺失Partial deletion of the first exon and 5' UTR region of ebony gene 钱远槐等, 1994;金珊等, 2005;金珊和曾庆韬, 2010;Freoa et al., 2023 注：表中AS-代表常染色体上的单基因隐性突变；AS+代表常染色体上的单基因显性突变；AS±为常染色体上的不完全显性（半显性）突变。“—”代表突变体暂无名称。Note: The symbol AS-represented the autosome recessive mutant with single gene, while AS+ represented the autosome dominant mutant with single gene, and AS± represented the incomplete dominant (semi-dominant) mutant. The symbol “—” represented no specific name for the mutant. -
图 1 黑色素合成信号通路（仿Noh et al.，2016; 高云等，2020）
注：绿色椭圆框代表色素合成通路的关键基因，黑色、黄色和白色框代表色素前体沉积到表皮的颜色。
Fig. 1 Pathway for Melanin synthesis in insects (Modified from Noh et al., 2016 and Gao et al., 2020)
Note: The green oval boxes represented the key genes of the pigment synthesis pathway, and the black, yellow and white boxes represented the color of the pigment precursor deposited on the epidermis, respectively.
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附表 1 昆虫黑化突变体及其形成机制与生物学特性变异
Appendix Table 1 Insect melanism mutants and the mechanisms and changing of biological characteristics
目
Order科
Family种名
Species名称及黑化阶段
Name and melanism stages遗传学规律
Genetic basis生物学特性变异
Changing of biological characteristics生理与分子机制
Physiological and molecular mechanism参考文献
References鳞翅目
Lepidoptera夜蛾科
Noctuidae棉铃虫
Helicoverpa armigera HübnerJBM
老熟蛹和成虫
Mature pupa and adultAS- 发育速度变慢, 成虫个体变小, 交配率降低, 生殖力下降
Decreased development rate, reduced adult body size, decreased mating rate and reproductive output酚氧化酶活力突增
Significantly increased activity of phenoloxidase能量代谢与细胞骨架组分蛋白的含量改变Changes of energy metabolism and cytoskeletal protein contents马伟华, 2005;Wang et al., 2005;Ma et al., 2008
何丽华等, 2010;何丽华等, 2012—
老熟蛹和成虫
Mature pupa and adultAS- 粘虫
Mythimna separate (Walker)—
成虫
AdultAS- 幼虫和蛹发育加快, 存活率提高; 产卵前期缩短且产卵量增加
Accelerated development of larva and pupa, increased survival, reduced pre-oviposition period and increased oviposition刘红兵和罗礼智, 2004;Jiang et al., 2007 甜菜夜蛾
Spodoptera exigua (Hübner)SEM
蛹
PupaAS± 发育速度加快, 蛹重增加, 交配时间延长, 生殖力提高
Accelerated development, increased pupal weight and mating period, increased reproductive capabilityTH与DDC过表达
Overexpression of TH and DDC刘子晶, 2006;刘向阳, 2007;刘思思, 2015;Liu et al., 2015a, 2015b 蚕蛾科
Bombicidae家蚕
Bombyx mori L.暗化型mln
幼虫、老熟蛹及成虫
Larva, mature pupa and adultAS- AANAT的功能失调及与儿茶酚胺代谢的交互作用
Dysfunction of AANAT and its interactive with catecholamineDai et al., 2010;Zhan et al., 2010;Qiao et al., 2012 浅暗化型Lightmelanism
幼虫、老熟蛹及成虫Larva, mature pupa and adultAS- 多巴、多巴胺含量改变 Changes of dopa and dopamine content 张彦等，2018，2021 黑蛾 Bm
成虫 Adult
soAS-Banno et al.，2005 幼虫、蛹及成虫, 以蛹最明显Larva, pupa and adult, but predominant in pupa ebony 基因开放阅读框的大片段缺 失 Large fragment deletion of the ORF of ebony Futahashi，2008 黑蛹bp
蛹PupaADC 基因起始位点调控序列插入转 座子 Insertion of transposons in the initial regulatory sequence of gene ADC Dai et al.，2015 从性黑蛾sml
雄性成虫Male adultAS-, 与性别关联Sex-related 寿命延长并具有 更高活力 Prolonged the lifespan and expressed higher activity yellow 与 laccase2 基因的上调表达 Up-regulated expression of gene yellow and laccase2 代方银等，2013；
He et al.，2016暗化蛾Melanism moth
幼虫及成虫Larva and adult mln-2AS- 茧重略有减轻 A slight decrease in weight of cocoon 杨明观等，2001 成虫Adult AS- 张凤林等，2009 黑缟(ps)
幼虫LarvaAS- 酪氨酸向多巴的转化速度提高 Accelerated transformation of tyrosine to dopa 刘丽丽等，2014；
张彦等，2017bd
幼虫LarvaAS- 雌性不育Infertility of female 表皮蛋白基因上调表达Up-regulated expression of cuticular protein gene Kawaguchi et al., 2007;Wu et al., 2016 天蛾科Sphingidae 烟草天蛾Manduca sexta L. bl
末龄幼虫Last-instar larva雌性连锁单基因, 并受其他基因修饰Female-linked single gene and modified by other genes 发育速率、体重、存活率及生殖力降低Decreased development rate, body weight, survival and fecundity 保幼激素含量不足, DDC基因上调表达Insufficient juvenile hormone content, and up-regulated expression of DDC Truman et al., 1973;Safranek and Riddiford, 1975; Hiruma and Riddiford, 2009 蛱蝶科Nymphalidae 眼斑蝶Bicyclus anynana (Butler) dl
幼虫LarvaAS- 生殖力降低Decreased fecundity Bear et al., 2010 赤拟谷盗Tribolium castaneum (Herbst) —
幼虫、蛹及成虫Larva, pupa and adultAS± 鞘翅的机械特性发生改变Changes of the mechanical characteristics of elytra 多巴胺含量增加Increased dopamine content Sokoloff et al., 1960;Kramer et al., 1984;Roseland et al., 1987;Lomakin et al., 2010 ebony
成虫AdultAS- 产卵量下降Decreased oviposition Park et al., 1945 鞘翅目Coleoptera 拟步甲科Tenebrionidae McGill black成虫Adult AS+ 发育时间延长Prolonged development time Stanley and Slatis, 1955;Edwards, 1958 杂拟谷盗Tribolium confusum Duval negret
老熟蛹及雄成虫Mature pupa and male adult雄性连锁的隐性调控Male-linked recessive regulation 发育速率变缓, 生殖力下降, 蛹期与成虫羽化期的存活率下降Slowdown of the development rate, decreased fecundity, decreased survival rates of the pupae and newly emerged adults Neelam and Bhatia, 1983 瓢甲科Coccinellidae 具斑食蚜瓢虫Coleomegilla maculata (DeGeer) 成虫鞘翅Elytra of adult AS± Allen, 2016 异色瓢虫 ml 羽化成虫畸形率降低; 鞘翅类胡萝卜素浓度升高; 抗紫外能力增强Decreased the malformation rate of newly emerged adults; Increased elytra carotenoid concentration; increased UV-resistance 李明凌等, 2022;李媛等, 2025;Sun et al., 2025 Harmonia axyridis (Pallas) 幼虫、蛹和成虫Larva, pupa and adult AS- 绿豆象Callosobruchus chinensis (L.) sC-b 繁殖力与生育力降低Decreased fecundity and fertility Kashiwagi and Utida, 1972 老熟蛹及成虫Mature pupa and adult AS- 豆象科
Bruchidae四纹豆象Callosobruchus maculatus (Fabricius) 成虫, 雌虫最明显Adult and predominant in female AS+ 雌虫个体大一些Increased body size of female adults Breitenbecher, 1921;Eady, 1991;Mano and Toquenaga, 2011 鹰嘴豆象Callosobruchus analis (Fabricius) 成虫
AdultAS- 适合度无影响
No significant effects on fitnessNumajiri et al., 2017 窃蠢科
Anobiidae烟草甲Lasioderma serricorne (Fabricius) 成虫
AdultAS- 交配行为与生活史均未发生改变No changes of mating behavior and life history Omae et al., 2012 黑檀体 ebony
成虫Adult交尾潜伏期增加, 交配及视觉行为异常Increased mating latent period, and exhibited abnormal mating and vision behavior β-丙氨酸代谢发生缺陷Defect in β-alanine metabolism Jacobsm, 1960;Hodgetts, 1972;Wittkopp et al., 2002;Kohn and Wittkopp, 2007 双翅目
Diptera果蝇科
Drosophilidae黑腹果蝇
Drosophila melanogaster Meigen黑体black
成虫Adult对视觉信号的反应异常Abnormal response to visible signals ADC基因突变Mutation of ADC gene
mdg4插入Yippee基因的5'端UTR区Insertion of mdg4 in 5' UTR region of Yippee geneHodgetts, 1972;Phillips et al., 2005 sable
成虫Adult√ Dean et al., 2022 黑条体ebonybsr
成虫Adult非孟德尔式遗传
Non-Mendelian inheritance求偶动作减少、体温升高Reduced courtship behavior, and increased body temperature ebony基因5'端UTR区和第1个外显子部分缺失Partial deletion of the first exon and 5' UTR region of ebony gene 钱远槐等, 1994;金珊等, 2005;金珊和曾庆韬, 2010;Freoa et al., 2023 注：表中AS-代表常染色体上的单基因隐性突变；AS+代表常染色体上的单基因显性突变；AS±为常染色体上的不完全显性（半显性）突变。“—”代表突变体暂无名称。Note: The symbol AS-represented the autosome recessive mutant with single gene, while AS+ represented the autosome dominant mutant with single gene, and AS± represented the incomplete dominant (semi-dominant) mutant. The symbol “—” represented no specific name for the mutant. -
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